of light by an object: an ordinary 60 W bulb emits about 100 cd. The amount of light received by an object per unit area is its illuminance, and is measured in lux. This unit is defined as the degree of illumination of a surface 1 m from a source of one candela radiating uniformly in all directions. Full sunlight may provide about 100 000 lux.
on how much they happen to be illuminated. A brightly illuminated patch of black on a white background still looks black, even though its luminance may be more than a neighbouring dimly illuminated patch of white. So adaptation has two functions: it enables you to cope with a very wide range of light levels, but it also provides information about albedo, the first step towards recognition.
Cones are found particularly in the middle of the visual field and provide very detailed information about the retinal image, being a little more than 2 µm in diameter, and are also responsive to colour. But cones have a high threshold, and can only function when the light is above some 10-2 cd/m2 (the photopic region). Below this, in the scotopic region, we’re forced to use the rods. At in-between levels we have an intermediate kind of vision called mesopic. The top of this mesopic region is at about 100 cd/m2, when the rods stop functioning because they’re completely saturated.
Rods are much more sensitive – in fact as sensitive as they could possibly be, since one individual rod can respond to a single photon of light – but the bad news is that in order to achieve this sensitivity they have to group themselves together into functional teams – numbered in thousands – by means of their neural connections in the retina. By pooling their information they enormously increase their sensitivity (whereas the random background noise tends to cancel out by being pooled, the signal one is trying to detect does not). But this is at the cost of throwing away a lot of information about the spatial detail of the retinal image, and also sacrificing the ability to distinguish wavelengths or colours; they are also slow, and they respond to a range of wavelengths that is slightly shifted in the blue direction – something called the Purkinje shift. The difference in the spectral sensitivity curves relates to the degree to which rods and cones absorb light of different wavelengths: the dashed green line shows how the absorption spectrum for the pigment in the rods, rhodopsin, has a very similar shape to the scotopic sensitivity.
Table 7.1 Vision under photopic and scotopic conditions | |||||||||||||||||||||
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ones less, and if you’re lucky they come to a point. To a first approximation, the way to get them to come to a point is to use a spherical surface. In the eye, there are three surfaces of this sort that act together to bring the images of distant objects to a focus on the retina: they are the cornea, and the front and back surfaces of the lens. The refractive index of the aqueous humour that separates the cornea and lens is much the same as that of the vitreous humour that fills the rest of the eye, and is about 1.34; that of the crystalline lens is only slightly greater than this, being about 1.41.
nature, since – being transparent – it obviously cannot have a blood supply. It obtains its nutrients and oxygen from the aqueous humour that bathes it on both sides, and is able to penetrate the lens because of fibrous nature. It is a fluid similar to plasma but with only some 1 per cent of its protein concentration and a peculiarly large amount of ascorbic acid. The aqueous humour is continuously secreted by the ciliary body, and passes through the iris into the anterior chamber where it eventually filters its way out into the canal of Schlemm, whence it ends up in the bloodstream. The resistance to its outflow generates an intraocular pressure of some 10-20 mmHg. Blockage may raise this pressure to the point where the flow of blood into the back of the eye is hindered, a serious condition called glaucoma which is a common cause of blindness.
called astigmatism. If, for example, it has a smaller radius of curvature in the horizontal plane than in the vertical, the far point when measured with a vertical line as test object will be closer than when a horizontal line is used. It is corrected with a cylindrical lens – in effect a section cut from a cylinder, just as a spherical lens is from a sphere: it focuses only in one meridian. Opticians test for astigmatism by means of a target like the one below, called an astigmatic fan; an astigmatic subject will see some of the lines more sharply than others, and this will tell the optician the angle at which a cylindrical lens should be placed in front of the eye to make the refractive power as nearly as possible equal in all meridians. The power of the cylindrical lens that is needed to do this, together with its meridional angle, make up the cylindrical correction that is the second part of a prescription for spectacles.
man-made lenses are nearly all spherical, simply because they’re easier to make, a spherical lens does not in fact bring rays to a point focus: they get bent too much as you go further out, and it is the resultant blur that is spherical aberration. The shape you really need is not a sphere but an ellipsoid. For surfaces that are small in comparison with their radii of curvature the difference is slight, and spherical aberrations are often negligible. But in the case of the eye, the aperture is of the same order of magnitude as the radius of curvature of the cornea, and the result is that rays entering near the periphery of the cornea are bent too much, and form a closer focus than those entering near the centre. To some extent Nature has compensated for spherical aberration, first of all by making a cornea that is not exactly spherical but tends towards the desired ellipsoid; and second, in that the refractive index of the lens is not constant throughout, but graded from a maximum of some 1.42 at its centre to about 1.39 at the edge, thus cancelling out, to some extent, the extra bending of peripheral light rays. The degrading effects of both spherical and chromatic aberration, and of other defects due to irregularities of the refracting surfaces, get worse as the pupil or aperture of the eye increases, and this in turn depends on the state of the iris: this is discussed on p. 137, in the context of visual acuity.
Box Table 7.1 | ||||||||||||||||||||||||||||||||||||||||||||
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the pointspread function has a diameter of about 1.5 arc min (measured half-way up); the worse your optics, the bigger this becomes.
contrast needed, and so the method provides a sensitive measure of acuity. The reason for the fall-off in contrast sensitivity at low frequencies is discussed later (p. 152).
the retinal image is blurred, one can still estimate where the peak is quite accurately. One can show that the longer the line the better one is, showing that accuracy is also being improved by averaging information over the whole line. Another pseudo-acuity task is the detection of stars, which may subtend extremely small angles at the eye. For instance, the bright star in Orion called Betelgeuse subtends only some 1/20 arc sec. But in a sense that figure is quite irrelevant: because of the pointspread, its image still has a width of 45 arc sec, and whether you detect it or not depends simply on how whether its luminance exceeds the absolute threshold, ΔI0.
conditions of dark adaptation the intrinsic acuity of the neural processing of the retinal image is so low that the poor optics contribute little to the overall blur, and the advantage of being able to increase retinal sensitivity by catching more light with a dilated pupil outweighs the disadvantage of slightly decreased acuity.
Table 7.2 Advantages and disadvantages of small and large pupils | |||||||
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all the difference in sensitivity between rods and cones (since the difference in threshold between one isolated rod and one cone is only about a log unit). But it is not so good for acuity. Because this neural pointspread is broader the further from the fovea, acuity is very much worse in the periphery than in the centre. And most important of all, as you dark-adapt, changing from central cone vision to rod vision with their enormous pooling of information, acuity drops off dramatically. If the contrast threshold as a function of spatial frequency is measured with a fixed pupil during progressive stages of dark adaptation, over 6 log units one finds a steady decrease in the cut-off frequency, the result of changes in the neural organization of the retina. One of the adaptational responses to reduced light levels, as we shall see, is an increase in the effective size of the ganglion cells’ summation pools, so that they can catch more light. But this obviously has the effect of increasing the degree of neural blur, and hence of reducing the overall acuity.
the consequence of what seems like a massive error of judgement on the part of Nature, namely the decision to have the retina inside out. The tips of the photoreceptors face outward, and they pass their information backwards through the retina: as a consequence, the nerve fibres from the retina find themselves inside the eye when they want to be outside. What they do is to come together to form the optic nerve, and crash their way to the exterior, together with the central retinal artery and vein, through a region called the optic disc, at about 15° to the nasal side of the optical axis. Since this area is consequently incapable of responding to light, subjectively it forms the blind spot. Although it is some 5° across, one is usually unaware of its existence because the brain tends to fill it in with whatever background colour or pattern immediately surrounds it. Close your left eye, and view the red cross in the figure below from a distance of about 30 cm: the alien will disappear, yet no discontinuity in the background will be apparent.
Table 7.3 Factors affecting visual acuity | |||||||||||||||||||||||||||||||||||||||||||||||||||||
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fovea is quite without rods, and the cones themselves are tightly packed to give the maximum information about image detail: the angular size of the rod-free region is about that of one’s fingernail with the hand fully extended. Cones in this region are about 2.3 µm across, corresponding to a visual angle of some half minute of arc. In the section of central monkey retina below, you can see that the depression arises because the retinal structures that elsewhere in the retina lie between the receptors and the lens – remembering again that the retina is inside-out in its layered structure – are here displaced to one side so as to cause the minimum scattering of incoming light. The supply of oxygen and nutrients for this region must derive almost entirely from the blood vessels that richly supply the choroid, the layer immediately superficial to the receptors and separated from them by the thin pigment epithelium, which helps to reduce scatter by absorbing the incident light. These features can all be seen in the section below, from a monkey retina.
or so, then breaking off and being destroyed. In the case of rods they seal themselves off near the bottom, to form a stack of flattened saccules or discs; in the cones they remain partially open: it is not obvious why. At the base of the outer segment the remains of the ciliary filaments and centrioles can be seen. The inner segment has mitochondria as well as the nucleus, and its inner end forms the synaptic junction with bipolar and horizontal cells. There is no doubt that the photopigment straddling the membranes of the outer segment discs plays a key role in transforming incident light into electrical changes, for if the pigment is isolated from the receptor it is found that its absorption of light of different wavelengths corresponds closely with the spectral sensitivity of the receptors themselves.
of regeneration. Estimates of the amount of pigment in the receptors of a living eye during particular stimulus conditions may be made by the technique of retinal reflection densitometry, in which one measures the amount and spectral composition of the light scattered back from the retina when a light is shone into the eye. In this way it is possible to track continuously the amount of rod or cone pigment in bleached form under relatively natural visual conditions. Alternatively, in microdensitometry, the spectral absorptions of individual receptors may be measured in a preparation on a microscope slide. As far as we know, the reactions that occur in rods and cones are fundamentally similar, though the regeneration of cone pigment is substantially quicker than in rods, so that under photopic conditions a smaller fraction of the cone pigment is in the bleached state than is the case for rods: this is one of the reasons why the cones are able to function at much higher light levels.